Size, shape and hue modulate attraction and landingresponses of the braconid parasitoid Fopius arisanus to fruitodour-baited visual targets

Jeanneth Perez • Julio C. Rojas • Pablo Montoya •

Pablo Liedo • Francisco J. Gonzalez • Alfredo Castillo

Received: 25 June 2011 / Accepted: 9 October 2011 / Published online: 16 November 2011! International Organization for Biological Control (IOBC) 2011

Abstract Female parasitoids are guided by multi-

sensory information, including chemical and physicalcues during host location. In the present study, we

investigated the behavioural responses of naıveFopiusarisanus (Sonan) females to visual targets baited withguava odour. In non-choice wind tunnel tests, the

attraction and landing responses of parasitoids to

spheres painted with different colours, and targets of

different shapes and sizes were evaluated. Femaleswere more frequently attracted and landed more often

on dark yellow targets than on targets with other

colours. There was no correlation between the bright-ness of each colour and the attraction or landing

responses. In contrast, both responses were correlatedwith relative reflectance (hue) of the coloured targets.

A positive correlation was observed between attraction

and hue, and a negative correlation between landingand hue. F. arisanus was attracted to and landed more

often on spheres than on other shape models. The

attraction response of this parasitoid was affected bythe size of the targets, with spheres of 10 and 12 cm

diameter being more attractive than spheres of 8, 6 and

4 cm diameter. The fact that F. arisanus females wereable to discriminate among visual targets that differ in

colour, shape and size stresses the importance of vision

during host location by this species.

Keywords Visual cues ! Spectral reflectance !Attraction ! Landing ! Host location ! Braconidae

Introduction

The host selection by parasitoids is a sequentialprocess that can be divided in host habitat location,

host location, and host acceptance (Vinson 1976;

Fatouros et al. 2008). At long-range, parasitoids foragebetween plants used by their host within their habitat,

Handling Editor: Torsten Meiners

J. Perez (&) ! J. C. Rojas ! P. Liedo ! A. CastilloEl Colegio de la Frontera Sur (ECOSUR), CarreteraAntiguo Aeropuerto km 2.5, 30700 Tapachula, Chiapas,Mexicoe-mail: ejperez@ecosur.mx

J. C. Rojase-mail: jrojas@ecosur.mx

P. Liedoe-mail: pliedo@ecosur.mx

A. Castilloe-mail: acastill@ecosur.mx

P. MontoyaSubdireccion de Desarrollo de Metodos, ProgramaMoscas de la Fruta DGSV SAGARPA, Central Poniente14, 30700 Tapachula, Chiapas, Mexicoe-mail: pablojmg17@hotmail.com

F. J. GonzalezCoordinacion para la Innovacion y la Aplicacion de laCiencia y la Tecnologıa, Universidad Autonoma de SanLuis Potosı, Sierra Leona 550, Lomas 2da. Seccion, 78210San Luis Potosı, SLP, Mexicoe-mail: javier@cactus.iico.uaslp.mx

123

BioControl (2012) 57:405–414

DOI 10.1007/s10526-011-9416-0

at medium-range they search for the hosts within hostplants, and finally at short-range females recognize

and accept their hosts (Volkl 2000). During the host

selection process, parasitoids are guided by multisen-sory information, including chemical, vibratory and

visual cues, which are used in an interactive manner.

Chemical cues seem to play an important role in thedifferent steps within the host selection process,

whereas visual cues are thought to be important only

after females have been stimulated by host odours(Vinson 1976).

The role of chemical cues during the host selection

process has been studied extensively in parasitoids,but the use of vision during this process is less

understood for this group of insects. Visual informa-

tion may consist of variables such as colour, shape,and size, among others. These parameters have

different behavioural relevance during host habitat

location at distance, and during host location after theparasitoid has landed on a plant. At long-range, colour

seems to be the most important visual parameter,

while at short-range, shape and size are crucial for hostrecognition and host acceptance (Wackers and Lewis

1999). Colour can be described in terms of three main

attributes: hue (dominant wavelength of reflectedlight), brightness (intensity of perceived reflected

light) and saturation (spectral purity of reflected light)

(Prokopy and Owens 1983). True colour vision ininsects has only been confirmed for a limit number of

orders, among them Hymenoptera (Chittka and Men-

zel 1992; Kelber et al. 2003). Most of hymenopteranspecies investigated so far have trichromatic vision

with UV (340 nm), blue (430 nm), and green

(535 nm) receptors (Peitsch et al. 1992). The capacityfor colour discrimination has been reported for several

species of parasitoids (Messing and Jang 1992;

Battaglia et al. 2000). For example, colour plays animportant role during host recognition and oviposition

by the parasitoid Aphidius ervi Haliday (Battaglia

et al. 2000). In addition to colour, other visualparameters may be used by parasitoids during host

location, including the size and shape (Vinson 1976).Host size and shape may be visual cues highly

distinguishable in almost any habitat because mor-

phological differences exist not only between plantsbut also between structures of the same plant (Wackers

and Lewis 1999). The use of these parameters during

host location has been documented for differentparasitoid species. For instance, Diachasmimorpha

longicaudata (Ashmead) females showed no prefer-ence for colour or shape, but displayed a clear

preference for large over smaller fruit-mimicking

models (Segura et al. 2007).Fopius arisanus (Sonan) is an egg-pupal endopar-

asitoid that attacks several species of tephritid fruit

flies, particularly those in the genus Bactrocera.Currently, F. arisanus is known to parasitize 21 fruit

fly species, and can develop in at least 18 species

(Quimio and Walter 2001; Rousse et al. 2005, 2006).The successful introductions of this parasitoid to

Hawaii and French Polynesia for the control of

Ceratitis capitata (Wiedemann) and Bactrocera dor-salis (Hendel), respectively, suggest that F. arisanushas potential for augmentative biological control of

fruit flies, alone or in combination with other controlmethods (Harris et al. 2000).

Host location behaviour of F. arisanus has been

described by Wang and Messing (2003). Chemicalcues play important roles during host location behav-

iour in this wasp (Liquido 1991; Altuzar et al. 2004;

Rousse et al. 2007a). However, there are contrastingreports regarding the role of visual preferences during

host location (Vargas et al. 1991; Cornelius et al. 1999;

Rousse et al. 2007b). With the aim of clarifying therole of visual preferences in host location by F.arisanus, this study was undertaken to investigate the

behavioural responses of female wasps to guavaodour-baited targets of different colours, sizes and

shapes in a laboratory wind tunnel.

Materials and methods

Insects

Parasitoids used in this study were obtained fromcolonies established at MOSCAFRUT mass-rearing

facility (SAGARPA-IICA) located in Metapa de

Domınguez, Chiapas, Mexico. This strain of parasit-oids was imported from Hawaii in 1998, where it was

reared on C. capitata eggs (Bautista et al. 1999).Parasitoids used for experiments had been reared from

Anastrepha ludens (Loew) eggs placed in pieces of

papaya at 22 ± 2"C, 60–80% RH, and 12:12 L:D hphotoperiod as previously described (Montoya et al.

2009). The eggs ofA. ludenswere obtained from adults

reared on artificial diet according to establishedprotocols (Domınguez et al. 2010). After emergence,

406 J. Perez et al.

123

parasitoids of both sexes were placed in wooden framecages 30 9 30 9 30 cm to allow mating. Adults were

given free access to honey and water dispensed on

cottonwool. One hour before the experiments, femaleswere placed individually in plastic containers (5 cm

high by 4 cm diameter) and allowed to acclimatize to

the wind tunnel room conditions. In all experiments,7–9 days old females were used. Parasitoids had no

prior contact with their hosts or host fruits.

Bioassays

The observations were performed in a wind tunnel(120 cm long 9 30 cm high 9 30 cm wide). The

wind tunnel was constructed using 1 cm thick clear

plexiglass. A fan was used to pull air, filtered byactivated charcoal, through the tunnel at a speed of

0.4 m s-1. Illumination was provided by four fluores-

cent tubes mounted 60 cm above the wind tunnelgiving a light intensity of 2380 lx. Green dots of

10 cm diameter made from cardboard were placed on

the tunnel floor to provide optomotor feedback for theparasitoids. Fruit odour consisted of five green guavas

(325.7 ± 13.2 g) (mean ± SD) without insect infes-

tation, placed inside a polyethylene bag that wassituated outside the wind tunnel. Filtered air

(1 l min-1) was passed through the bag and the odour

was conducted into the wind tunnel by means of tygontubing (0.4 cm internal diameter). The end of the tube

was connected to a hole (0.7 cm) in the centre of the

visual target from which odour emanated at a flow rateof 0.6 l min-1. The target stimulus was hung in the

centre of the wind tunnel, 16 cm from the upwind end.

The distance between platform release and the targetstimulus was about 1 m.

Each observation started by placing the plastic

container with one female on a 12 cm high platform atthe downwind end of the tunnel. The parasitoid was

released and observed for 5 min. Female behaviour

was recorded, specifically the zigzagging upwindflight towards the target stimulus and landing on the

target after such upwind flight. If the parasitoid did nottake off or did not show zigzagging upwind flight after

5 min, the test was stopped and the insect was

considered to be a non-responding individual. Here-after, the term attraction is used to indicate such

zigzagging upwind flights. Parasitoids were used

once. In all experiments, the responses of parasitoidsto the visual targets were evaluated in non-choice

tests. All bioassays were conducted between 08:00 hand 14:00 h at 24 ± 2"C, 65 ± 5% RH.

Response to models of different colours

The responses of F. arisanus females to styrofoamspheres of different colours baited with guava odour

were evaluated. Preliminary observations showed thatfemales were not attracted to spheres without the

odour. Spheres of 8 cm diameter were painted with the

following colours of acrylic paint (Comex Group,Mexico): white, black, brown, red, blue, light green,

dark green, light yellow and dark yellow. The spectral

reflectance curves of the different colours were mea-sured using a spectrometer (USB4000-VIS-NIR,

Ocean Optics, Dunedin, FL, USA) with an optical

resolution of*1.5 nm (full width at half maximum), atungsten-halogen light source (LS-1, Ocean Optics,

Dunedin, FL) and a reflection probe (R200-7-VIS-NIR,

Ocean Optics, Dunedin, FL, USA). The raw reflectancespectra were corrected for detector dark current and

normalized to the spectrum obtained from the light

source reflected on a white reference standard (Fig. 1).Five replicates per colour were made daily in a random

order and the experiment was repeated on different

days until completing the 45 replications per colour.

Response to models of different shape

We evaluated the responses of F. arisanus females tomodels of different shapes with similar surface area

Wavelength (nm)400 500 600 700

Spec

tral

ref

lect

ance

(%)

0

3

6

20

40

60

80W

Dy

R

LyB

DgBrLg

Bl

Fig. 1 Spectral reflectance curves of the nine colours used inthe experimentDy dark yellow, Ly light yellow, B blue,Wwhite,Br brown, Bl black, R red, Lg light green and Dg dark green

Responses of the braconid 407

123

(313 cm2). All models were made using styrofoamand were painted dark yellow. This colour was chosen

because it was the most attractive to F. arisanusfemales in the previous experiment. The modelsevaluated had either three-dimensional (sphere, cube,

pyramid, and ellipse) or flat shapes (circle, square,

rectangle, and triangle). Five replicates per shape wereperformed daily in a random order and the experiment

was repeated on different days until completing 45

replicates per shape.

Response to models of different size

The responses of F. arisanus females to dark yellow

spheres of 14, 12, 10, 8, 6, and 4 cm diameter were

tested in this experiment. The size of spheres werechosen on the basis of previous studies with this

species and considering the range of size of fruits used

by F. arisanus hosts. Five replicates per size weremade daily in a random order and the experiment was

repeated on different days until completing 45 repli-

cates per size.

Statistical analysis

We used a log likelihood ratio contingency-table

analysis to test whether responses (attraction or

landing) differed among treatments. Fisher’s exacttest was used when counts were low. When significant

effects were found, odds ratios were used to make

comparisons between treatments.In the first experiment, numbers of attraction and

landing responses were correlated with the brightness

and hues of visual targets, using methods describedelsewhere (Katsoyannos and Kouloussis 2001).

According to this procedure whether the insect

response to coloured target depends only on bright-ness, a strong correlation between the frequency of

responses and total brightness was expected. If the

insect response depends on the specific wavelengths(hues), then the response should correlate with relative

reflectance values of the colours. Relative reflectancewas considered to be the amount of light reflected by a

colour within a given wavelength interval relative to

the total amount of light reflected by this colour withinthe range of the spectrum considered.

The total reflectance values of the colours used in this

study were obtained as percentages of the total amountof light reflected over a white reference sphere between

350 and 750 nm. Reflectance values were: white =100, dark yellow = 51, light yellow = 14.1, dark

green = 7.9, light green = 9.7, blue = 9, brown =

8.9, black = 7.1, and red = 24. Relative reflectancewere calculated for every colour as the amount of light

reflected by each successive 10 nm wide wavelength

interval, in relation to the total amount of light reflectedby that colour between 350 and 750 nm.

Results

Response to models of different colours

Colour influenced the attraction of F. arisanus females

towards the spheres baited with fruit odour (G = 21.1,df = 8, P = 0.007). Females were more attracted to

dark yellow models than to light yellow, blue, black,

red, light green, and dark green models (Fig. 2a).There were no differences in the numbers of females

that were attracted to dark yellow, white, and brown

targets. The landing behaviour of parasitoids on thetargets was also influenced by colour (G = 67.44,

df = 8, P\ 0.001). Females landed more often on the

dark yellow, light yellow, brown, black, red, and lightgreen models than on blue, white and dark green

targets (Fig. 2b).

There was no correlation between the total amountof reflected light of each colour and attraction

(r = 0.44) or landing (r = -0.399) responses (P[0.05). In contrast, both responses were correlated withrelative reflectance (hues) of the coloured targets. A

positive correlation was detected between attraction

and hue, and a negative correlation between landingand hue (Fig. 3). Females were more attracted to hues

reflecting maximally between 550 and 620 nm (dark

yellow), with a maximum peak at 600 nm (r = 0.795).Parasitoids landed less often on hues reflecting

maximally between 420 and 510 nm (blue, light

green), with a minimal peak at 490 nm (r = -0.943).

Response to models of different shape

The shape of visual targets affected attraction

(G = 24.2, df = 7, P = 0.001) and landing (G =19.0, df = 7, P = 0.008) responses of F. arisanus.Parasitoids were more attracted to spheres than to

square, rectangle, or pyramid models. Attraction ofF. arisanus females to spheres was not significantly

408 J. Perez et al.

123

different from that of circle, ellipse, cube and triangle

models (Fig. 4a). Parasitoids landed more often on

sphere, circle, and cube models than on the trianglemodel. The landing response on ellipse, square,

rectangle and pyramid were intermediate and not

significantly different from that on sphere, circle andcube or triangle (Fig. 4b).

Response to models of different size

The size of the visual targets affected attraction of

F. arisanus females (G = 33.7, df = 5, P\ 0.001).In contrast, the landing response was not affected by

the size of the models tested (Fisher test, P = 0.76).

Parasitoids were more attracted to spheres of 10 and

12 cm diameter than to spheres of 4, 6 and 8 cmdiameter. The attraction of F. arisanus to spheres of

14 cm diameter was intermediate and not significantly

different from that to spheres of 10 and 12 cmdiameter (Fig. 5).

Discussion

We showed that some visual cues in combination withfruit odour might play a role in the host location

0

20

40

60

80

100

DarkYellow

LightYellow Green

DarkGreen

Att

ract

ion

resp

onse

(%)

Colour

bc

a

bc bcbc

bc

c

ab ab

a

0

20

40

60

80

100

DarkYellow

LightYellow

Blue White Brown Black Red Light

Blue White Brown Black Red LightGreen

DarkGreen

Lan

ding

res

pons

e (%

)

Colour

a

a a

aa

a

b

b

b

b

Fig. 2 Attraction (i.e.zigzag upwind flighttowards the visual targetstimulus) (a) and landing(b) (±S.E.) of F. arisanusfemales on visual targets ofdifferent colour. Barscapped with the same letterare not significant different(P[ 0.05)

Responses of the braconid 409

123

behaviour of naıve F. arisanus females. We found thatfemales of this parasitoid were attracted and landed

more often on yellow spheres compared with spheres

of other colours. Previously, Vargas et al. (1991)reported that yellow and orange traps captured more

F. arisanus than those caught by blue and black traps.

Rousse et al. (2007b) showed that naıve F. arisanusfemales landed less often on yellow and white spheres

than on the black and dark green spheres. These

authors concluded that F. arisanus females respondmainly to achromatic cues. However, the response that

we obtained towards the dark yellow and brown

spheres showed that they were equally attractive toF. arisanus, although they differ widely in the amount

of reflected light, suggesting that for this species the

hue is more important than brightness. An explanationfor differences between both studies may be due to the

experimental protocols. In our case, experiments were

performed using fruit odour in combination withvisual targets, and this may have affected the parasit-

oid responses. Thus, we consider that the response of

F. arisanus females to odour-baited visual targets maybe related to host foraging, while the responses of the

parasitoids reported by Rousse et al. (2007b) may only

imply the ability of females to detect and respond tovisual targets, without direct link to the host location

process. On other hand, our bioassays were performed

in a wind tunnel with controlled illumination, such thatthe amount of light received by the coloured visual

targets of the different treatments was constant

throughout the whole experiment. Rousse et al.

(2007b) performed experiments in field cages wherethe amount of light received by each visual target

likely varied depending on their position relative to the

sun and the target, and variation in solar and clima-tological condition, thus influencing the parasitoid’s

responses. Indeed, the received colour of an object

depends upon the interaction between the ambientlight and the reflectance colour of the object. Thus, an

object may have a different appearance in each

environment (Endler 1993). Another reason is that inour study the visual targets were not placed against a

particular background (black, white, gray). Consider-

ing that insects likely perceive foliage as gray, thusagainst this achromatic background, any different

wavelength other than that of foliage will produce at

least some colour contrast (Chittka et al. 1994).The preference for the colour yellow we found in

F. arisanus has been reported for other hymenopterans,

including egg parasitoids (Lobdell et al. 2005). In thecase of fruit fly parasitoids, Diachasmimorpha juglan-dis (Muesebeck) chose yellow walnut fruits over black

fruits despite that the latter are more likely to containhost larvae (Henneman 1998). In field cage tests,

D. longicaudata females discriminated between vari-

ous spheres painted with different colours with yellowas the most attractive colour (Messing and Jang 1992).

In the same way, several studies have shown that

yellow colour is highly attractive to different species offruit flies (Katsoyannos 1987; Vargas et al. 1991; Drew

et al. 2003; Pinero et al. 2006; Lopez-Guillen et al.

2009). For example, C. capitata females were mainlyattracted to yellow, followed by red, orange, black, and

green spheres, whereaswhite and blue sphereswere the

least attractive (Katsoyannos 1987).The shape and size of targets are other visual

parameters that can be used during host location by

parasitoids. The hue and brightness of visual targetscan be perceived before insect landing, whereas the

shape and size gain in importance at lesser distances.

Vinson (1976) pointed out that size and shape areusually secondary stimuli during foraging, and

become important during host acceptance. However,in some cases these stimuli seem to be crucial during

host location. For instance, Henneman et al. (2002)

suggested that at distance,D. juglandis females tend toignore the odour and use visual cues, because the

odour plume is turbulent and difficult to locate the host

fruit. We have found that F. arisanus females showedbetter responses to spherical than to rectangular or

-1.0

-0.8

-0.6

-0.4

-0.2

0.0

0.2

0.4

0.6

0.8

1.0

350 390 430 480 520 560 600 650 690 740

Cor

rela

tion

coe

ffic

ient

(r)

Wavelength (nm)

Attraction Landing

P < 0.05

P < 0.05

Fig. 3 Correlation coefficient between the responses (attrac-tion and landing) of F. arisanus females to targets of ninedifferent colours, and the relative reflectance values of eachcolour at successive 10 nm wavelength, from 350 to 750 nm.Statistical differences are indicated above or below of the dash-lines

410 J. Perez et al.

123

square models. It is important to mention that we

performed this bioassay using yellow models only.Therefore, further experiments are needed to investi-

gate whether shape may interact with colour as it is

documented for other insects (Bernays and Chapman1994). Also, we found that parasitoids were more

attracted to spheres between 10 and 14 cm diameter.

Diachasmimorpha longicaudata females also showedpreferences for larger spheres (Segura et al. 2007).

Similarly, the shape and size of visual targets influence

host selection in different species of fruit flies. Adultsof C. capitata and Anastrepha fraterculus (Wiede-

mann) are more attracted to spherical forms than to

rectangular models (Cytrynowicz et al. 1982). Lopez-

Guillen et al. (2009) reported that spherical modelscaptured more Anastrepha obliqua (Macquart) adults

than those caught by rectangular, triangular, or

circular models. Also, they observed that spheres of8, 10, 12 cm diameter captured more A. obliqua adultsthan smaller spheres. It has been suggested that fruit

flies are more attracted to spherical models becausethis shape mimics the fruits used by insects for

oviposition (Katsoyannos 1989). A similar idea may

be applied to explain the results obtained withF. arisanus females, where colour, size and shape

could have a significant influence in host location.

0

20

40

60

80

100

Att

ract

ion

resp

onse

(%)

Shape

a

abab

abc

d

cd

abc

bcd

a

0

20

40

60

80

100

Sphere Circle Ellipse Cube Square Rectangle Triangle Pyramid

Sphere Circle Ellipse Cube Square Rectangle Triangle Pyramid

Lan

ding

res

pons

e (%

)

Shape

b

ab

a

ab

ab

ab

bbb

Fig. 4 Attraction (i.e.zigzag upwind flighttowards the visual targetstimulus) (a) and landing(b) (±S.E.) ofF. arisanus onvisual targets of differentshape. Bars capped with thesame letter are notsignificant different(P[ 0.05)

Responses of the braconid 411

123

Insects usually use multisensory information duringforaging (Harris and Foster 1995). In this way, Hebets

and Papaj (2005) proposed several hypotheses con-

cerning the role of multimodal signals, including thepossibility that a combination of cues facilitates the

detection of a target at different distances, where the

detection of one sensory modality alerts the insect tothe presence of the other. In this sense, we used fruit

odour in all our experiments because parasitoids were

not attracted to visual targets when they were offeredalone. Our results emphasize the importance of the

interaction of different cues during host selection by

this parasitoid. Thus, although we used the sameolfactory cue, females were able to discriminate

among the different models offered indicating the

importance of visual cues at long-range. The impor-tance of visual cues at short-range was also observed

in this study. For example, although the white colour

was as attractive as the dark yellow, only 36.6% ofparasitoids that were attracted to white spheres landed

on them. The remaining females that did not land on

white spheres (23%) hovered at 1–2 cm from thetarget for about 3 min, and then returned to the

downwind end of the tunnel, sometimes landing on the

release platform. This result is in agreement with theidea that vision has a critical role in the final stages of

host location, partly because the lack of odour

gradients makes it most improbable that odour wouldguide an insect directly to a plant, but also because a

flying insect needs to use vision during landing(Bernays and Chapman 1994). The multimodal sen-

sory integration has been reported for other parasitoid

species as well. Jang et al. (2000) suggest that during

foraging, visual and olfactory cues, both alone and incombination, affect the behaviour of D. longicaudatafemales. Likewise, Jonsson et al. (2005) reported an

increase preference for flower odour when colourswere added in the bioassays performed with the

ichneumonid females of Phradis interstitialis Thom-

son and Tersilochus heterocerus Thomson, parasitoidsof Meligethes spp. (Coleoptera: Nitidulidae). Finally,the tachinid parasitoid Exorista japonica Townsend

uses both visual and olfactory cues to locate the hosthabitat of the noctuid pest Mythimna separata(Walker) (Ichiki et al. 2011).

Our results suggest that F. arisanus attraction tocoloured targets is due to the hue rather than the

intensity of reflected light. We have showed that

F. arisanus can discriminate between colours, butwithout odours these visual cues mean nothing for

naıve F. arisanus females, at least within the limits of

our bioassays. From this, we infer that females areinitially stimulated by fruit odours and are then guided

to targets by a combination of visual and chemical

cues. The fact that parasitoids were able to discrim-inate among visual targets that differed in colour,

shape and size, stresses the importance of vision

during host location by this species.

Acknowledgments We are grateful for the technical assistanceof Florida Lopez (PROGRAMA MOSCAFRUT) and ArmandoVirgen (ECOSUR). Rosalba Morales provided logistic support.Many thanks go to Javier Valle Mora for statistical advice andFrancisco Infante and Trevor Williams for a critical review of aprevious version of themanuscript. J. Perez gives special thanks toEl Consejo Nacional de Ciencia y Tecnologıa (CONACYT) ofMexico for the doctoral scholarship.

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100

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Att

ract

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aab

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Fig. 5 Attraction (i.e. zigzag upwind flight towards the visualtarget stimulus) (±S.E.) of F. arisanus on visual targets ofdifferent size. Bars capped with the same letter are notsignificant different (P[ 0.05)

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Author Biographies

Jeanneth Perez Received her MSc degree from El Colegio dela Frontera Sur, Mexico. At present she is a Ph. D candidate atthe same institution. Her research interests are chemicalecology, insect behaviour and biological control of tropicalinsects.

Julio C. Rojas Received his PhD from Oxford University in1998. He is a research entomologist at the Department ofTropical Entomology, El Colegio de la Frontera Sur, Mexico.His interests include the behaviour and the chemical ecology ofherbivorous and carnivorous insects.

Pablo Montoya Doctor in Sciences from the UniversidadNacional Autonoma de Mexico in 1999. He is a Sub-Director

and researcher in the Moscafrut Program SAGARPA-IICA ofMexico. He has experience in the management of fruit flies,particularly in biological control, detection of populations, andthe application of the Sterile Insect Technique.

Pablo Liedo Ph. D. in Entomology, University of California -Davis (1989). He is a research entomologist at the Departmentof Tropical Entomology, El Colegio de la Frontera Sur,Mexico. His area of interest is insect ecology and pest control,with particular emphasis on biodemography, population ecol-ogy, biological control and the Sterile Insect Technique.Research work has been focused on fruit flies.

Francisco J. Gonzalez Ph.D. in electrical engineering from theSchool of Optics and Photonics, University of Central Florida,Orlando, in 2003. He is currently a Professor at the UniversidadAutonoma de San Luis Potosı, Mexico. His research interestsare the application of physics, optics and engineering in non-invasive medical diagnosis.

Alfredo Castillo Doctor in Science from El Colegio dePosgraduados in 2005. He is an entomologist at the Departmentof Tropical Entomology, El Colegio de la Frontera Sur,Mexico. He conducts research on ecology and biology oftropical insects.

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